A Checklist of the Birds of Taiwan
Tzung-Su Ding
Abstract
A checklist of the birds of Taiwan is a basic and essential part for studies on the avifauna of Taiwan. The latest checklist proposed by Chinese Wild Bird Federation (CWBF) in 1995 took a serious attitude on examining and verifying new records proposed after World War II, but regrettably it is not updated with current changes on the nomenclature of some taxons. In this article, a brief review on the history of ornithology of Taiwan and current classification systems of birds is provided. Also the CWBF1995 checklist is examined and its conflicts with Howard and Moore (1991), the classification system which CWBF (1995) claimed to follow, are reported. An alternative checklist of the birds of Taiwan that is based on the species account in CWBF1995 but follows the SAM (Sibley-Ahlquist-Monroe) classification system is prepared. This new checklist will serve as basis of the other parts of my dissertation.
History of ornithology of Taiwan
Taiwan, an island off the coast of Southeast China, is predominately populated by Chinese who started to immigrate in the sixteen century. Before the colonization of Chinese, The aborigines are undoubtedly the first group of people who discover birds species and own a great extent of knowledge on the birds of Taiwan. However, the indigenous peoples did not have written language to bear their knowledge. The first written record mentioning the birds in Taiwan is from the book, Encyclopedia of Taiwan <Tai-Wan-Fu-Chi>, published by officials in 1696 (Lin 1997). Before Taiwan annexed by Japan in 1896, there were more than 17 similar books published for revision or local counties in Taiwan (Lin 1997). It is a tradition of Chinese to write official book on a certain county or province about its local history, geography, culture, and natural history. However, this kind of books hardly sets up any foundation for ornithology of Taiwan. The authors are basically literary men who did not go to field for collection; the records, mainly based on hearsay or imagination, are short and sometimes mythological; and the names of birds are usually copied from ancient literature and not recognizable (Swinhoe, Horikawa <cited in Lin 1997>, Lin 1997). The modern ornithology of Taiwan was started by occidental natural historians after the opium war (1840-1842), when several Chinese ports were opened for foreign trade and residence.
The first scientific record of birds of Taiwan is from an exploring expedition of U.S. Navy (Lin 1997). On its route passing by Taiwan, on the 25th of September, 1854, William Stimpson, the accompanied scientist, made a small collection of three bird species in Taiwan and later examined and published by John Cassin (Cassin 1862). However, in spite of the historical significance of Stimpson's collection, which accidentally started the history of Taiwan's ornithology, the scientific study of Taiwan ornithology is now unanimously accepted to have established by the extensive work of Robert Swinhoe, who arrived Taiwan in 1856. Robert Swinhoe, a British diplomatist born in Calcutta in 1836 and educated in London, entered the consular service as interpreter in 1854, served as vice-consul and consul at various ports of China during the following twenty years, and died in London in 1877, at the age of 41 (Hachisuka and Udagawa 1950). His first expedition to Taiwan was made in 1856, reaching Heongsan and journeying around the coast for two weeks (Swinhoe 1859a). Later in 1858 he joined a trip circumnavigating Taiwan (Swinhoe 1859b) and in 1860 he was appointed as vice-consul in Taiwan, the first foreign resident diplomatist in Taiwan. During his stay from 1860 to 1866, he vigorously explored all types of territory and extensively collected specimens of birds, mammals, reptiles, amphibians, fishes, insects, shells, and plants. Many of his discoveries of Taiwan's birds were described in consecutive acticles appeared in the magazine, "Ibis" (Swinhoe 1860, 1862a, 1862b, 1863a, 1863b, 1864a, 18654b, 1865a, 1865b, 1866a, 1866b, 1866c, 1869, 1872, 1877). Swinhoe is the first one who ever published the checklists of birds of Taiwan (Swinhoe 1863a) and of China (Swinhoe 1863c). 227 species (including five endemic species) out of the 460 bird species recorded in Taiwan now were solely discovered by him (Lin 1997).
In the October of 1873, the exact month that Swinhoe left China for health consideration, Professor J. B. Steere of the University of Michigan arrived Takao (now Kaoshyong) and spent five months in Taiwan (Hachisuka and Udagawa 1950). He collected a novel endemic species, Liocichla steeri, which was named later by Swinhoe (Swinhoe 1877). After this event, for twenty years there was almost no publication concerning the avifauna of Taiwan, until A. P. Holst made his expedition to Taiwan in 1893. Holst stayed in Taiwan for more than one year, ascended Mt. Yushan, and collected many zoological specimens, including an endemic tit species that bears his name, Parus holsti (Seebohm 1894, 1895). J. D. D.La Touche, another outstanding zoologist in China, also visited Taiwan three times from 1893 to 1895 (La Touche 1895, 1898).
In 1896, after the Sino-Japanese war, Taiwan was annexed by Japan. Japanese zoologists gradually took over the task of discovery and research. In 1897, Tsunesuke Tada, at the request of the Tokyo Imperial University, explored many places that had not visited by western zoologists, especially Botel Tobago (now Lanyu) (Hachisuka and Udagawa 1950). In 1906, Water Goodfellow stayed in Taiwan for three months and successfully collected many novelties at high altitudes (Ogilvie-Grant 1906). His collection was later examined by W. R. Ogilvie-Grant and included all the 6 endemic species of Taiwan that had not been discovered by that time (Ogilvie-Grant 1906). One of them is Mikado Pheasant, of its striking beauty of plumage drew the attention among the British ornithologists. A. Owston sent Yonetaro Kikuchi, a Japanese collector, to Taiwan for alive Mikado Pheasants and other novelties. Owston kept sending fresh materials collected by Kikuchi to L. W. Rothschild, E. J. Hartert, Ogilvie-Grant, and C. Ingram and contributed valuably to Taiwan's ornithology (Rothschild 1907, Ogilvie-Grant and La Touche 1907, Rothschild and Hartert 1907, Hartert 1909, Ingram 1909, Ingram 1910). In 1907, Ogilvie-Grant and La Touche compiled a complete account of Taiwan's birds (Ogilvie-Grant and La Touche 1907). The number of bird species knew to be occurred in Taiwan became 260, an increase of 73 on Swinhoe's 187 species enumerated in 1863 (Swinhoe 1863a). In 1912, Goodfellow made his second expedition of Taiwan but discovered no new species (Ogilvie-Grant 1912). After the death of Owston in 1915, except of three or four articles, the European interest in Taiwan's ornithology was ceased and it was abruptly taken over by Japanese ornithologists.
During the Japanese occupation period from 1896 to 1945, except of the years in Pacific War, Taiwan was the southmost territory of Japan and its tropical fauna provided Japanese a novel ground of exploration. The birds from Taiwan were popular in the gardens and pet shops of Japan (Hachisuka and Udagawa 1950). Taiwan is also convenient for Japanese ornithologists collecting or taking trips because it is near to Japan and was under the Japanese control. In spite of its colonial purpose, the Japanese government in Taiwan took a serious attitude for establishing academic institutions in Taiwan. The first museum in Taiwan, State Educational Museum (destroyed during World War II), was found in Tainan in 1902. The second one, Taihoku Museum (now Taiwan Museum), was found in Taipei in 1908. Both played an important role in collecting and discovering new species in Taiwan. The first university in Taiwan, Taihoku Imperial University (now National Taiwan University), found in 1928 with Zoology Department from its initial stage, although later impeded by wars, was also found for advancing the academic research in Taiwan.
Most of the prominent Japanese ornithologists of the Japanese occupation period visited Taiwan and published articles or books on the birds of Taiwan (Hachisuka and Udagawa 1950). Among them, Seinosuke Uchida and Nagmichi Kuroda are the Japanese ornithologists who greatly contributed to the ornithology of Taiwan at the early stage (Hachisuka and Udagawa 1950). Uchida compiled a list of the birds of Taiwan (Uchida 1912), adding 30 species to the former list (Ogilvie-Grant and La Touche 1907). In 1915, Uchida prepared a very useful book, "The Birds of Japan", and its third volume is entirely devoted to the birds of Taiwan, 301 species known at that time (Uchida 1915). In 1916, Kuroda, a well-known bird taxonomist, took a field trip in Taiwan, examined the specimens stored in the museums of Taipei and Tainan, and added 34 new species to Taiwan's avifauna (Kuroda 1916). Kuroda also concluded that the avifauna of Lanyu have a stronger affinity to Philippine rather than to Taiwan, and proposed to elongate the "Neo-Wallace line" to divide Lanyu and Taiwan (Kuroda 1929), a treatment still widely accepted by current zoogeographers. During his career, lots of Taiwan new records were examined and published by him (e.g. Kuroda 1920a, 1920b, 1920c, 1922, 1927, 1928a, 1928b, 1930, 1931, 1936) (Hachisuka and Udagawa 1950). Yoshimaro Yamashina made a trip to Taiwan himself and sent his collectors to Taiwan from 1932 to 1936 and collected over 2000 specimens of birds and a great amount of nests and eggs (Yamashina 1937, Yamashina and Yamada 1937, 1938). Till now, our knowledge on the breeding behavior of birds in Taiwan are still largely through Yamashina's studies. Tadao Kano, a devoted researcher, extensively explored the Tsugitaka (now Sheishan) mountains and published a book on the fauna of Taiwan's mountains (Kano 1940), detailing the ecological and altitudinal distribution of most of the resident bird species.
During the period of Japanese occupation, although all the endemic species had been discovered by occidental ornithologists, Japanese discovered the rest of resident species (Lin 1997) and enumerated the birds of Taiwan from 260 species in 1907 (Ogilvie-Grant and La Touche 1907) to 393 species in 1942 (Hachisuka et al. 1942). Japanese ornithologists published several books and more than 160 articles entirely or partly devoting to the ornithology of Taiwan by 1945 (most of them documented in Hachisuka and Udagawa 1950). Their efforts extended the scope of Taiwan's ornithology from solely discovering species and taxonomical studies to ecological and ethological studies. Masauji Hachisuka integrated the studies done by Japanese and, along with Tatsuo Udagawa, prepared a detailed and useful review on the ornithology of Taiwan (Hachisuka and Udagawa 1950, 1951), making a perfect period of the Japanese era.
Right after World War II, civil war broke out in China and the Nationalist (KMT) government fled to Taiwan in 1949. Most of the Japanese in Taiwan had been sent back to Japan and virtually there was no Taiwanese ornithologist existing. No Chinese ornithologist came to Taiwan and the harsh political atmosphere was also not favorable for scientific research in fundamental disciplines. The ornithology of Taiwan was totally stagnated. In the first book written in Chinese on Taiwan's fauna, "The Vertebrates of Taiwan", (Chen 1956), the chapter on Class Aves is completely compiled by the works formerly done by occidental and Japanese ornithologists. The chapter on bird in the second edition (Chen 1969) is identical to the first edition, still with no original work from Chinese or Taiwanese. This dark was not broken until the first light shed from the studies conducted by U.S. Army and American ornithologists during the 1960's. The "Migratory Animal Pathological Survey" (MAPS) was conducted by 13 teams in 9 countries of East Asia from 1963 to 1971 (McClure 1998). Totally 1,165,288 birds of 1218 species were banded by this project funded mainly by US Army (McClure 1998). The team in Taiwan was led by Drs. Paul Alexander and Sheldon Servinghaus of Tunghai University in Taichung (McClure 1998). The team members trained by this project served as seeds to the rejuvenation of Taiwan's ornithology. From 1972 to 1975, the "Ecological Survey of Forest Birds of Taiwan", the first research ever funded by Taiwan government, was solely conducted by Taiwanese ornithologists (Chen and Yen 1977). Since then, ornithological studies on Taiwan are rapidly growing and all led by Taiwanese. Right now, in the universities and research institutes of Taiwan there are more than one dozen of researchers with a Ph.D. degree entirely or partially devoting their research to Taiwan's ornithology. Besides, the activity of bird watching is getting popular in Taiwan. The Taipei Wild Bird Society was found in 1973, Taichung in 1975, Kaoshyung in 1979 (Lin 1997). Till 1999, there are more than 17 such kind of societies in Taiwan. Those societies nourish a great number of enthusiastic amateur ornithologists and provide vast energy to promote various aspects in academic research and conservation movement.
Classification Systems of the Birds of the World
One essential objective of zoology is to reconstruct a detailed history of the evolution of animals. Classification, the attempt to assign animals to the most appropriate taxonomic group, is to try to portray the evolutionary relationship of animals. However, the bird fossil records, the most persuasive evidence, are incomplete and fragmentary; ornithologists have to reply on other characters to hypothesize the evolutionary relationship of groups, so-called avian systematics. As long as the avian systematics advences, the avian classification changes. After Darwin's evolution theory, the current theoretical foundation for systematics and classification, there has been a long succession of classification system. Before the second half of twentieth century, the characteristics in morphology and comparative anatomy were the only materials for constructing classification system. But these kinds of characters are often problematic because some similarities are due to adaptative convergences. Recently, more attention has been given to ethological characteristics, such as songs, courtship display, and to other physiological and biochemical aspects, including the degree of similarity of proteins and DNA. The monumental work by James L. Peters, the 15 volumes of "Checklist of the Birds of the World" (Peters 1934-1987), is the most prevailing classification system in the twentieth century. Most of the other classification systems (e.g. Merony et al. 1975, Clements 1981, Howard and Moore 1980, 1991) are similar to Peter's system except of some new discoveries and taxonomical revisions. However, their systems are fundamentally based on morphological and anatomical comparison and subsequently have quietly weak support at the macrosystematic level because of the lack of objective evidence.
Since the 1980s, the field of avian systematics has been stirring by a profound impact from Charles Sibley and his colleagues. They employed the DNA-DNA hybridization techniques, enabling the direct comparison of genetic materials of two species, and proposed a new and revolutionary classification system (Sibley and Monroe 1990, Monroe and Sibley 1993) based on their results from the DNA-DNA hybridization of some 1702 bird species (Sibley and Ahlquist 1990). Most treatments in their classification system, so-called SAM (Sibley-Ahlquist-Monroe) system, corroborate many conclusions that have been previously resulted from traditional anatomical approach (Paine 1998). However, some other conclusions at macrosystematic level are pretty novel and have been attracting wild debates and controversies (e.g. Mayr and Bock 1994). The traditional classifications are familiar to most ornithologists but lack of strong and convincing evidence on its macrosystematic treatment. The new biochemical approach is unanimously accepted as valuable and promising for avian systematics but its current conclusion has been criticized as "full of uncertainty" and "subject to further testing".
For the purpose of dessertation, I select the SAM system as the base of nomenclature and classification. The first and most important reason is that, at microsystematic level (species and subspecies), which is still vastly relied on morphological, ethological, and zoogeographical characteristics, SAM system is more updated than any other tradictional classification systems (e.g. Merony et al. 1975, Clements 1981, Howard and Moore 1980, 1991). Most of the newly published books, although some do not agree with the treatment of SAM system at macrosystematic level (order, family), are closer to SAM system in the nomenclature of species. Second, the differences at macrosystematic level among those classification systems are not related to the purposes of my dissertation and will not affect the results. Last, except that the unfamiliarity or confusion critisms may still hold, the other main criticisms such as "full of flaws" or "subject to verification" are counter, or even more, applicable to the tradictional classification systems.
Current checklists of the birds of Taiwan
After the monumental review of Hachisuka and Udagawa (1950, 1951), several checklists on the birds of Taiwan have been prepared (e.g. Yen 1979, Chang 1985, Wang 1991, CWBF 1995). The latest one, the checklist proposed by Chinese Wild Bird Federation (CWBF) in 1995, is the outcome of several meetings that cautiously examined and verified the new records that have been proposed since the end of World War II. That checklist is the most complete and reliable account of Taiwan's bird species so far. However, regrettably some species on that list continue to use traditional nomenclature. Since the era of Japanese occupation, except of some controversial cases, many bird species have been unanimously split or lumped. The CWBF1995 checklist claimed it follow the classification system of Howard and Monroe (1991) but aparently it fails to update some taxonomical changes that have even been accepted by Howard and Monroe (1991).
The first part of the rest of article is a report on the taxonomical and nomenclatural inconsistency among the Howard and Moore 1991 and CWBF1995 checklist. The second part is another checklist of Taiwan's birds that species account is based on CWBF1995 but classification and nomenclature follow SAM system (Monroe and Sibey 1993) (MS1993). The impact of adapting SAM system to the checklist of Taiwan's birds is also briefly introduced.
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